Biogeography and Ecological Distribution of Shallow-Water Benthic Foraminifera from the Auckland and Campbell Islands, Subantarctic Southwest Paciﬁc

Journal of Micropalaeontology, 26: 127–143. 0262-821X/07 $15.00 2007 The Micropalaeontological Society Biogeography and ecological distribution of shallow-water benthic foraminifera from the Auckland and Campbell Islands, subantarctic southwest Paciﬁc

BRUCE W. HAYWARD, HUGH R. GRENFELL, ASHWAQ T. SABAA & RHIANNON DAYMOND-KING Geomarine Research, 49 Swainston Rd, St Johns, Auckland, New Zealand (e-mail: [email protected])

ABSTRACT – One hundred and forty-eight species of benthic foraminifera are recorded from depths shallower than 80 m around the subantarctic Auckland (130 spp.) and Campbell (71 spp.) Islands, southwest Pacific. Comparisons with other circum-polar, subantarctic island groups suggest that they all have relatively low diversity, shallow-water benthic, foraminiferal faunas, with their sheltered harbours dominated by species of Elphidium, Notorotalia, Cassidulina, Haynesina and Nonionella-Nonionellina. More exposed environments are dominated by a small number of species of Cibicides, Miliolinella, Rosalina, Quinqueloculina and Glabratellidae. The extremely low species richness (three species) in high-tidal grass-dominated salt marsh on Campbell Island is similar to that reported from Tierra del Fuego at a similar latitude. The faunas of Auckland and Campbell Islands have their strongest affinities (70–75% species in common) with New Zealand’s three main islands, 460–700 km away. Ten percent of their fauna has not been recorded from mainland New Zealand, reflecting one endemic species and a small element of apparently subantarctic and bipolar-restricted species. Since there have been no shallow-water (<500 m) links to other lands since these two Miocene volcanic islands were formed, it is concluded that most benthic foraminiferal species have arrived in suspension in eddies of surface water, many since the peak of the Last Glacial. J. Micropalaeontol. 26(2): 127–143, October 2007.

KEYWORDS: subantarctic islands, Auckland Islands, Campbell Island, shallow-water benthic foraminifera, biogeography

INTRODUCTION Previous work and source of samples studied The Auckland and Campbell Islands groups were first discov- Location and setting ered by Europeans (fur sealers) in 1806 and 1810, respectively The uninhabited Auckland (50( 45# S, 166( E) and Campbell (McNab, 1909) and, since then, many scientific expeditions have (52( 30# S, 169( E) Islands are the southern-most land areas visited them and collected shallow-marine biological samples within New Zealand’s Exclusive Economic Zone in the (summarized in Chilton, 1909; Hayward & Morley, 2005). From southern SW Pacific Ocean (Fig. 1). Auckland Islands the late 1870s until the early twentieth century, New Zealand (626 km2) and Campbell Island (113 km2) are separated from government steamers made regular visits to both island groups New Zealand’s South Island by 460 km and 700 km, respect- to check for castaways. Natural scientists sometimes joined ively (Peat, 2003). Both are the partly eroded remnants of these trips and their collections and observations added to Miocene basalt shield volcanoes sitting on uplifted basement knowledge of the subantarctic biota (e.g. Hutton, 1879). On one and oceanic sedimentary rocks (Cook, 1981) that form of these trips in 1897, several samples of seafloor sediment were seamounts sitting above the 500 m background average depth dredged from depths shallower than 20 m near Enderby Island of the Campbell Plateau. (Auckland Islands) and Campbell Island. These samples were During the Pleistocene, glaciers flowed down radiating stream washed and floated by New Zealand amateur microscopist R. L. valleys on both island groups and eroded out wide ‘U-shaped’ Mestayer, who made numerous slides of qualitative picks of profiles. Most of these valleys were flooded by rising sea-level foraminifera (F201105, F201106, F201107, F201108), mostly during each interglacial, forming long, deep (up to 70 m) inlets unilocular forms (now held in the collections of the Institute of and harbours (Fig. 1). The Auckland Islands have a coastal Geological and Nuclear Sciences, Lower Hutt). fringe of low rata (Metrosideros) forest that passes upwards into The only previous work devoted entirely to the foraminifera dense scrub. Campbell Island has Dracophyllum scrub and of New Zealand’s Subantarctic Islands has been by Australian grassland extending down to the shore (Peat, 2003). The average micropalaeontologist, Frederick Chapman (1909). He provided rainfall on both islands is 1000–1500 cm, with precipitation extensive documentation of the species’ composition (168 taxa) averaging 325 days each year on Campbell Island (Peat, 2003). of five sediment samples dredged during the 1907 scientific Much of this rain soaks into the peaty soil beneath the forest or expedition by the Philosophical Institute of Canterbury scrub and flows out through the intertidal zone as acidic, (Chilton, 1909). Four of the samples were from outer-shelf tannin-stained seepage. depths (100–170 m) offshore from Auckland, Enderby, Snares Mean annual sea surface temperature range is 7–11(Catthe and Bounty Islands and one sample was from sheltered harbour Auckland Islands and 1.5–2(C cooler at Campbell Island. Both waters (16 m, Perseverance Harbour, Campbell Island). island groups are today surrounded by Subantarctic Surface Chapman described and named six new species or varieties Water within weaker parts of the northeast-flowing Sub- (Table 1), all of which came from the deeper samples off antarctic Current (Fig. 1), between the Subtropical Front and Enderby and the Snares Islands. He also identified ten species in Subantarctic Front (Carter et al., 1998). the shallow-water sample from Campbell Island (Table 2).

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Fig. 1. Location of the Auckland and Campbell Islands southeast of New Zealand, also showing surface current directions and oceanic fronts. Sample locations around Auckland and Campbell Islands. SAF, Subantarctic Front; STF, Subtropical Front.

Marjorie (M. K.) Mestayer, conchologist at the Dominion small dredge samples of sediment from Carnley Harbour. These Museum (Wellington), obtained sediment from c. 50 m depth at samples, and slides of foraminifera picked from them, were the Auckland Islands in the 1910s and donated some with its deposited in the collections of the New Zealand Geological molluscs to Baden (A. W. B.) Powell at Auckland Museum Survey (where Fleming was employed as a palaeontologist; (L13513, F201015). In the 1940s, several natural scientists were F201102, F201162) and Auckland Museum (L6184, L13574). In stationed on the Auckland Islands to watch for enemy warships his study of New Zealand ostracods (including the Auckland (Peat, 2003) and, during their time there, Charles (C. A.) Islands), Norcott Hornibrook (1953) used M. K. Mestayer, Fleming, Bill (W. H.) Dawbin and Jack (J. H.) Sorenson all took Fleming and Sorenson material (above), and Paul Vella (1957)

128 Benthic foraminifera of Auckland and Campbell Islands

Original name Author, year Type location Name accepted in this study

Miliolina chrysostoma Chapman, 1909 off Enderby Island, Auckland Islands Triloculina chrysostoma Planispirinina antarctica Chapman, 1909 off Enderby Island, Auckland Islands Lagena lagenoides var. nuda Chapman, 1909 off Enderby Island, Auckland Islands Lagena quadrata var. carinata Chapman, 1909 off Snares Islands Lagena enderbiensis Chapman, 1909 off Enderby Island, Auckland Islands Spirillina novaezealandiae Chapman, 1909 off Snares Islands Spirillina novaezealandiae Notorotalia aucklandica Vella, 1957 Carnley Harbour, Auckland Islands Notorotalia aucklandica Nonion flemingi Vella, 1957 Carnley Harbour, Auckland Islands Nonionellina flemingi Elphidiononion simplex aoteanum Vella, 1957 Carnley Harbour, Auckland Islands Haynesina depressula Siphotextularia mestayerae Vella, 1957 Auckland Islands Siphotextularia mestayerae Textularia subantarctica Vella, 1957 Auckland Islands Textularia subantarctica

Table 1. Species of foraminifera described as new from material collected around New Zealand’s Subantarctic Islands.

Original name and authority Recorded by Location Name accepted in this study

Rhizammina indivisa Chapman, 1909 Campbell Island Rhizammina algaeformis Reophax scorpiurus Chapman, 1909 Campbell Island Reophax subfusiformis Reophax nodulosa Chapman, 1909 Campbell Island Reophax nodulosa Uvigerina angulosa Chapman, 1909 Campbell Island Trifarina angulosa Sphaeroidina bulloides Chapman, 1909 Campbell Island Sphaeroidina bulloides Truncatulina variabilis Chapman, 1909 Campbell Island Cibicides dispars Anomalina coronata Chapman, 1909 Campbell Island Discanomalina coronata Rotalia clathrata Chapman, 1909 Campbell Island Notorotalia aucklandica Nonionina boueana Chapman, 1909 Campbell Island Nonionellina ﬂemingi Polystomella macella Chapman, 1909 Campbell Island Elphidium advenum limbatum

Table 2. Species of foraminifera previously recorded from inner shelf depths (0–50 m) around the Auckland and Campbell Islands. described five new species of foraminifera from the Fleming and et al., 1982) and thus do not accurately represent the true living M. K. Mestayer samples (Table 1). Many of the old qualitative biodiversity near the islands. Because of this, the unilocular foraminiferal slides from R. L. Mestayer, M. K. Mestayer, records have been removed from species lists before making Fleming and Dawbin have been examined during this study, comparisons between regions. with additional taxa, especially unilocular, being added to the species list (Appendix A). MATERIAL AND METHODS Recently, the deep-water foraminiferal faunas (deeper than 50 m) have been described and mapped for the Campbell Samples Plateau and surrounding region, including mid- and outer- Most sediment samples used in this study (Fig. 1, Appendix B) shelf depths (>50 m) around Auckland and Campbell Islands were taken by BWH during a visit to the two island groups in (Hayward et al., 2007). March, 2004. Subtidal seafloor samples were taken by a hand- hauled surficial grab or small box dredge, which tended to Objectives sample the upper 5–7 cm of surface sediment. Intertidally a The main objective of this study is to document more fully the 10 cm3 sample of the upper 1 cm of surface sediment was taken foraminiferal biodiversity of these distant subantarctic islands, with a plastic tube. This set of samples was supplemented by compare their faunal composition with that of mainland New older samples held in the collections of the Auckland War Zealand (Hayward et al., 1999) and other islands in the sub- Memorial Museum, and the Institute of Geological and Nuclear antarctic zone around the world (e.g. Heron-Allen & Earland, Sciences, Lower Hutt (above). Some of these samples consisted 1932; Earland, 1933, 1935; Parr, 1950). From these studies it is entirely of a rich qualitative pick of foraminifera, but, where hoped to provide a better idea of foraminiferal biogeography in washed bulk sediment was also present, a new quantitative pick the SW Pacific and Southern Ocean and infer modes of disper- of the foraminiferal fauna was made to produce the total species sal. list (Appendix A). There was no opportunity to distinguish living from dead shells through staining. Unilocular species Faunal slides (Appendix B) are housed in the collections of One complicating factor in this study is the highly variable the University of Auckland, Auckland War Memorial Museum richness and diversity of unilocular lagenid species that some- and Institute of Geological and Nuclear Sciences, Lower Hutt times occur in huge numbers in shallow-water subantarctic (prefixed by AU, L, and F20, respectively). Census data are samples. It is suspected that many of these small, low density available digitally as Appendix C (available online at http:// species may have been winnowed out of deeper-water sediments www.geolsoc.org.uk/SUP18275. A hard copy can be obtained and transported into the shallows in suspension (e.g. Murray from the Geological Society Library).

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Laboratory processing Samples were washed gently over a 63 µm sieve, and the sand residue dried and microsplit down to an amount containing approximately 200 benthic specimens. All benthic foraminifera were picked from the microsplit, mounted on faunal slides, identified and counted (Appendix C). All taxa were identified to species level. Specimens of planktic foraminifera were counted during picking. Some of the samples from the harbours that were rich in small (63–150 µm) specimens which may have been transported in, were dry sieved over a 150 µm sieve and additional picks of this coarse fraction were made (samples with ‘c’ suffix).

Statistical analyses and diversity measures Quantitative samples (>63 µm) selected for statistical analysis Fig. 2. Number of species of shallow-water (<100 m) benthic foraminifera (with and without unilocular taxa) from around northeast North, comprised 16 from Auckland Islands and 21 from Campbell South, Stewart, Chatham, Snares, Auckland and Campbell Islands. Island – seven Campbell and two Auckland Islands samples were intertidal. Three Auckland and 12 Campbell Islands coarse rich unilocular fauna present in several historic collections samples (>150 µm) were also included. Thus the quantitative (unilocular species from Auckland Islands=31; from Campbell faunal data consist of census counts of 93 foraminiferal species Island=7). Among the more common New Zealand-wide species from 52 samples. The data matrix was transformed by convert- that are present at Auckland Islands but absent from the more ing counts to percentages of sample totals. A Q-mode cluster distant Campbell Island are Scherochorella moniliforme, Tro- analysis dendrogram classification was produced using a Chord chammina inflata, Patellina corrugata, Rosalina vitrevoluta and ffi dissimilarity coe cient matrix. The mathematical definition of four species of Quinqueloculina. Within the harbours the most ffi this coe cient is given in Sneath & Sokal (1973). noteworthy difference is the abundance of Nonionellina flemingi Detrended canonical correspondence analysis (DCCA, Ter and, to a lesser degree, Haynesina depressula at Auckland Braak, 1985) was used to summarize the percentage faunal data Islands, and their near absence from Campbell Island. The in a two-dimensional ordination plot and relate them to a set of harbours in both groups have almost identical diversity, with measured environmental and diversity factors (Appendix B). 61–63 species each in the quantitative samples and, apart from Three indices of benthic foraminiferal species diversity the two species already mentioned, they share the same suite of (Appendix B) were determined (Hayek & Buzas, 1997): (1) dominant species. Lack of sampling of salt marsh or brackish Fisher Alpha Index, (number of species standardized by environments in the Auckland Islands prevents comparisons number of individuals counted); (2) Information Function, H (a between the agglutinated faunas, and the lack of sampling of combination of the number of species present and, to a lesser high energy, shallow-water environments at Campbell Island extent, the evenness of species counts); and (3) Evenness, E (a prevents comparison between faunas characteristic of these. measure of dominance versus evenness of species counts).

TAXONOMIC COMPOSITION OF THE SUBANTARCTIC Taxa restricted to the Subantarctic in the New Zealand region FAUNA Here we record fourteen species (other than unilocular) that have not previously been recorded from elsewhere in the New Species richness Zealand region – eight from the Auckland Islands, two from A total of 148 species of benthic foraminifera, including uni- Campbell Island and four from both. There are several ad- locular species, are recorded here from the Auckland (130 spp.) ditional species (Discanomalina coronata, Lagenammina difflugi- and Campbell (71 spp.) Islands (Fig. 2). If unilocular species are formis, Rhizammina algaeformis, Siphogenerina dimorpha) that excluded, there are 113 species from these two groups (Auckland have been recorded occasionally from bathyal and abyssal – 96 spp., Campbell – 64 spp.). This number of species (without depths elsewhere in the region but are here recorded from uniloculars) is of a similar order to the 53 spp. recorded from shallower than 80 m at both island groups. A similar phenom- around Snares Islands (Hayward et al., 1999) and the 102 spp. enon is known from the shelf around Antarctica, where a from the Chatham Islands (Fig. 2; Hayward & Grenfell, 1999). number of species occur that are restricted to deeper water in As happens elsewhere in the world (Gibson & Buzas, 1973; lower latitudes (Mikhalevich, 2004). Murray, 1991) there is a decrease in total benthic foraminiferal Haplophragmoides manilaensis – this large agglutinated species species richness in shallow water, from warm to cool through the with inflated chambers (Pl. 1, figs 1–2) occurs widely in salt 18( of latitude of the New Zealand region (Fig. 2). marshes around the world. It was described from subtropical conditions in the Gulf of Mexico, but has also been recorded Comparisons between Auckland and Campbell Islands from subantarctic marshes as far south as Tierra del Fuego The main difference between their benthic foraminifera is the (Scott et al., 1990). greater diversity recorded from Auckland Islands (130 spp.) Labrospira spiculolega –This distinctive species (Pl. 1, figs 3–4) compared with Campbell Island (71 spp.), partly as a result of was described from 120–160 m depth, south of Tasmania the greater variety of habitats sampled and partly because of the (Parr, 1950) and is here recorded from inner-shelf depths at

130 Benthic foraminifera of Auckland and Campbell Islands

Plate 1. Explanation of Plate 1. Some common and distinctive benthic foraminifera from southwest Pacific subantarctic islands. Scale bar 0.1 mm. figs 1–2. Haplophragmoides manilaensis Andersen, 1953. BWH163/24, AU18127, Campbell Island intertidal MHWN. figs 3–4. Labrospira spiculolega (Parr, 1950). BWH 169/11, F201106, Auckland Islands 12 m. figs 5–6. Spiroplectammina biformis (Parker & Jones, 1865). BWH 168/14, AU18124, Campbell Island intertidal MHW. Apertural view is slightly oblique. figs 7–8. Verneuilinulla advena (Cushman, 1922). BWH168/22, F201106, Auckland Islands 12 m. figs 9–10. Patellinoides conica Heron-Allen & Earland, 1932. BWH169/12, F202511, Auckland Islands 80 m. Peripheral view is slightly oblique. fig. 11. Cerobertina tenuis (Chapman & Parr, 1937). BWH 169/15, F201106, Auckland Islands 12 m. fig. 12. Cerobertina tenuis (Chapman & Parr, 1937). BWH 169/16, F201106, Auckland Islands 12 m. fig. 13. Elphidium advenum f. limbatum (Chapman, 1907). FP4320, F201106, Auckland Islands 12 m. fig. 14. Heronallenia lingulata (Burrows & Holland, 1895). BWH 125/17, F202072, Snares Islands 100 m. figs 15–16. Heronallenia lingulata (Burrows & Holland, 1895). BWH 125/16, F202072, Snares Islands 100 m. fig. 17. Notorotalia aucklandica Vella, 1957. Topotype BWH 131/8, F201002, Auckland Islands 12 m. figs 18–19. Notorotalia aucklandica Vella, 1957. Topotype BWH 131/8, F201002, Auckland Islands 12 m.

the Auckland Islands. It may be endemic to shelf depths in the King George Island, South Shetland Islands (Gaz´dzicki subantarctic region of the SW Pacific. & Majewski, 2003; Majewski, 2005) in the Southern Spiroplectammina biformis (Pl. 1, figs 5–6) – appears to be a Hemisphere. widespread, shallow-water, higher-latitude, Northern Verneuilinulla advena (Pl. 1, figs 7–8) – this small agglutinated Hemisphere species (Murray, 1991) with a few rare records species appears to occur in shallow sheltered environments at from higher latitudes in the Southern Hemisphere. Its type high latitudes in both hemispheres, with its types from 10– locality is in 100 m of water off Greenland (Parker & Jones, 40 m in Hudson Bay, Canada (Cushman, 1922). It has been 1865) and it is widespread around the Gulf of St Lawrence recorded from abyssal depths at low latitudes (Loeblich & (e.g. Schafer & Cole, 1982) and in Scandinavian fiords (e.g. Tappan, 1994). Alve & Nagy, 1986; Hanslik & Nordberg, 2006). It also occurs Spirillina obconica – this uncommon species has been recorded in higher latitudes in the North Pacific (Murray, 1991) and at from off northern Europe and Florida in the Northern

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Hemisphere and from 100–300 m depth off the subantarctic Prince Edward and Kerguelen Islands (Brady, 1884). Laryngosigma williamsoni – this species was described from off the coast of the UK (Terquem, 1878) and has been recorded from a number of European localities. Neolingulina viejoensis – this rare, distinctive species has been recorded previously from 40–50 m depth off Peru and Korea (McCulloch, 1977). It probably is more widespread and under-reported. Pseudolingulina bradii – an uncommon but distinctive species, described from Indonesia, and also recorded from Hawaii, and 80–1000 m in the South Atlantic (Brady, 1884). Cerobertina tenuis (Pl. 1, figs 11–12) – this appears to be a southern, high-latitude species, with its type locality in upper abyssal depths (2600 m) between Australia and the Auckland Islands (Chapman & Parr, 1937). Bolivina translucens – this is a cosmopolitan species that is often identified under several other names. Fursenkoina cf. riggi – this small distinctive and uncommon species may be restricted to the subantarctic, having been described from the Gulf of San Gorge, southern Argentina (Boltovskoy, 1954) and recorded from Isla de los Estados (Thompson, 1978). Notorotalia aucklandica (Pl. 1, figs 17–19) – appears to be endemic to the Auckland and Campbell Islands. It is morpho- logically most similar to several New Zealand species of Notorotalia from which it presumably evolved (Vella, 1957). Patellinoides conica (Pl. 1, figs 9–10) – this species occurs in mid- Fig. 3. Number of species in selected genera and families of shallow- water (<100 m) benthic foraminifera from around northeast North, to high latitudes in the North and South Atlantic, and was South, Stewart, Chatham, Snares, Auckland and Campbell Islands. described from the subantarctic region, near the Falkland Islands (e.g. Heron-Allen & Earland, 1932). northeast North Island but only four occur in the subantarctic Heronallenia lingulata – around New Zealand, this small species (Fig. 3) – mostly absent are species of Pileolina. The same (Pl. 1, figs 14–16) is restricted to subantarctic Auckland and pattern also occurs in the shallow-water Rosalinidae and the Campbell Islands plus the Snares Islands. It appears to have a genus Quinqueloculina (Fig. 3). Other notable absences from the scattered cosmopolitan distribution in deeper water, but subantarctic are New Zealand’s most common foraminiferal occurs in relatively shallow water of higher latitudes in the SW genus in sheltered harbour settings (Hayward et al., 1999), Pacific and southern Australia (e.g. Brady, 1884). A similar Ammonia, and several of the most abundant open-marine distribution is displayed by H. unguiculata which occurs in species, Gaudryina convexa, Patellinella inconspicua and Tro- shallow water at the Auckland and Campbell Islands but at chulina dimidiatus. depths >200 m off northern New Zealand (Heron-Allen & Adifferent pattern emerges in the shallow-water members of Earland, 1922) and northeast Australia (Sidebottom, 1918). the Rosalinidae, with similar levels of species richness (13–14 spp.) off northeast North Island and in the subantarctic (Fig. 3). New Zealand taxa absent from the subantarctic. Notable by their absence from New Zealand’s subantarctic islands are a number Affinities with shallow-water foraminifera of the New Zealand of species that are common or abundant in shallow water region around mainland New Zealand (Hayward et al., 1999). These Auckland and Campbell Islands have 68–76% of their benthic include a number of the brackish and high tidal agglutinated foraminifera (excluding uniloculars) in common with the South species (e.g. Ammobaculites exiguus, Ammoscalaria tenuimargo, Island and northeast North Island (Table 3). In contrast, a Ammotium fragile, Jadammina macrescens, Miliammina obliqua, smaller percentage (52–59%) live around the nearer, but smaller, Trochamminita salsa), with 13–14 species occurring around the Chatham and Stewart Islands and just 31–32% have been two main islands and a combined total of nine at the sub- recorded around the Snares Islands (Table 3). An explanation antarctic islands (Fig. 3). The abundant shallow-water genus for the low levels of co-occurrence with the Snares is probably Elphidium is represented by only one species at Auckland and the lack of sheltered, harbour or brackish environments there. Campbell Islands and two additional species at the Snares, Explanations for the lower level of co-occurrence with Stewart whereas nine species are present at the South Island and 11 off and Chatham Islands are more difficult. It may reflect the lower northeast North Island (Fig. 3). In the related genus Notorotalia, number of samples and environments examined around these five species occur around the main islands, but only two in two island groups. the subantarctic (including the subantarctic endemic N. Of most significance is the observation that 94–98% of the aucklandica). Similarly, nine species of Glabratellidae occur off Snares fauna co-occurs around the South and North Islands.

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Campbell Auckland Snares Stewart Chatham South NE North Total no. Island Islands Islands Island Islands Island Island of spp. Campbell Island 100 73 32 59 57 68 71 64 Auckland Islands 47 100 31 55 52 74 76 96 Snares Islands 38 55 100 75 74 94 98 53 Subantarctic Islands 47 71 40 53 49 74 77 134

Table 3. Percentage of the benthic foraminiferal fauna (excluding unilocular spp.) of subantarctic Campbell, Auckland and Snares Islands (separately and jointly) in common with other parts of the New Zealand region.

This reflects the fact that only one Snares species (Heronallenia Ross). These associations are dominated by varying combina- lingulata) is restricted to the subantarctic islands, whereas the tions and abundances of four species. Association 4 (Nonionel- more distant Auckland and Campbell Islands have 13 species lina flemingi) occurs in muddy sand at 17–42 m depth in Tagua not found further north in the New Zealand region. Bay and Camp Cove (Figs 1, 5) in the cleaner, shallower parts of Carnley Harbour, Auckland Islands. Association 7 (Cassidulina FAUNAL ASSOCIATIONS carinata–Nonionellina flemingi) is most widespread inside the From the cluster analysis dendrogram we identified 10 faunal sheltered deeper (42–80 m), muddier parts of the long Carnley, associations (Fig. 4, Table 4). Five associations (1–3, 5, 8) are from Campbell Island and six from the Auckland Islands (4, 6–10), with only one association (8) occurring in both places (Fig. 5). With three exceptions, the clustering separated the coarse faunas (>150 µm) from the total sand range faunas (>63 µm), with the coarse faunas from Auckland Islands (assoc. 6) separated from the Campbell Island ones (assoc. 5). The two total faunas (A3, C11) that cluster with the coarse faunas have minimal fine fraction specimens present, and the same is true for the two total faunas (A5, A8) that cluster with the coarse fauna (A6c) forming association 4.

Agglutinated benthic foraminiferal associations The ﬁrst-order subdivision in the dendrogram separates faunas that are dominated by agglutinated species and live in Tucker Cove, Campbell Island (assocs 1–3), from all the remaining faunas, which are dominated by calcareous species. Two of the three agglutinated associations (1, 2) occur in high tidal salt meadow composed entirely of the salt-tolerant grass Isolepis cernua. The Haplophragmoides-dominated association (2) occurs in a 20 cm wide (tidal elevation) zone around extreme high water spring (EHWS) with the Miliammina fusca-dominated association (1) present between EHWS and mean high tide level. These two high tidal associations have the lowest diversity (=0.5–0.6, H=0.2–0.9) but highest level of evenness (E=0.5–0.8) of all associations. The third agglutinated association (3) is dominated by Eggerelloides scaber and Textularia earlandi, with secondary Paratrochammina bartrami and Spiroplectammina biformis (Fig. 6). This also occurs in Tucker Cove (Fig. 5), but in anoxic organic-rich mud at 4 m depth in the middle of the narrow bay. It has moderate levels of species diversity and evenness (Table 4).

Calcareous benthic foraminiferal associations A second-order subdivision within the calcareous faunas separates associations 4–8 from 9–10 (Fig. 4). The first of these two groups of associations is restricted in occurrence to the muddy sand and sandy mud on the floor of the relatively sheltered, Fig. 4. Dendrogram classifications of shallow-marine foraminiferal elongate harbours of Campbell (Perseverance Harbour) and sample associations produced by cluster analysis using Chord distance. Auckland Islands (Carnley, Norman, Smith Harbours, Port Associations 1–10 were selected by inspection of the dendrogram.

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Ass Dominant species Depth (m) n H E Islands

1 Miliammina fusca EHWS 3 0.5 0.2 0.50 Campbell 2 Haplophragmoides manilaensis, H. wilberti EHWS 3 0.6 0.9 0.80 Campbell 3 Eggerelloides scaber, Textularia earlandi 4 1 2.3 1.5 0.49 Campbell 4 Nonionellina ﬂemingi, Notorotalia aucklandica 17–42 3 1.9 1.2 0.40 Auckland 5 Elphidium advenum MT–40 13 2.4 1.0 0.36 Campbell 6 Notorotalia aucklandica, Elphidium advenum 10–17 3 2.9 1.6 0.44 Auckland 7 Cassidulina carinata, Nonionellina ﬂemingi 42–80 7 6.9 2.3 0.44 Auckland 8 Cassidulina carinata 4–40 14 4.1 1.5 0.29 Auckland, Campbell 9 Pileolina radiata 9 1 2.5 1.0 0.24 Auckland 10 Rosalina irregularis 0–8 3 4.4 2.0 0.48 Auckland

, Fisher Alpha Index (number of species standardized by number of individuals counted); H, Information Function (a combination of the number of species present and, to a lesser extent, the evenness of species counts); E, Evenness (a measure of dominance versus evenness of species counts); EHWS, extreme high water spring; MT, mid-tide. Table 4. Dominant species, depth range, number of stations (n), mean of diversity indices (, H, E) and island distribution of foraminiferal sample associations (Ass).

Fig. 5. Distribution of faunal associations produced by cluster analysis (1–10, Fig. 2) in the shallow water around Auckland and Campbell Islands.

Norman and Smith Harbours in the Auckland Islands. This around the Auckland Islands. Association 9 (Pileolina radiata)is association has subdominant Sigmoilopsis elliptica, Trifarina known from one sample at 9 m deep from just inside the western angulosa and Eilohedra vitrea and has the highest species diver- entrance to Carnley Harbour (Fig. 5) where big swells and sity in the study (=6.9, H=2.3). Association 6 (Notorotalia strong currents sweep over the seafloor. Subdominant are aucklandica) is limited to the slightly more exposed, shallower Quinqueloculina suborbicularis and Notorotalia aucklandica. It (10–17 m) Port Ross, in the northern Auckland Islands. has relatively low species diversity and the lowest evenness of all Association 5 (Elphidium advenum) is restricted to Campbell the associations (E=0.24) as a result of the strong dominance by Island and consists of all the coarse faunas throughout P. radiata. Association 10 (Rosalina irregularis) occurs in three Perseverance Harbour, at depths of 4–40 m, plus the total fauna samples at 0–8 m depth from either side of Enderby Island (C11) from mid-tidal level on the muddy sand beach in Tucker (Fig. 5). The samples from the north side (A15–A16) are not in Cove (Fig. 5). Association 8 (Cassidulina carinata) contains all situ and comprise sand washed up onto the rocks by storms, but the total faunas (>63 µm) from the same samples as association are considered representative of the fauna in the shallow sub- 5, plus the total faunas from two samples (A2, A4) at 10–18 m tidal just offshore. Subdominant in this association are P. depth in Port Ross, Auckland Islands. The main difference radiata, Miliolinella subrotundata and Quinqueloculina incisa between these two associations is in the relative abundance of (Fig. 6). small Cassidulina, which comprises 40–70% of the total faunas and less than 2% of the coarse faunas and, to a lesser extent, the DISCUSSION greater abundance of small Eilohedra vitrea in the total faunas. Distribution of benthic foraminiferal associations (Fig. 5) The second of the two calcareous groups occurs in cleaner The DCCA ordination (Fig. 7) shows that there are four sands from more exposed, wave- or current-swept locations separate groups of associations.

134 Benthic foraminifera of Auckland and Campbell Islands

Fig. 6. Mean relative abundance and standard deviation of the characterizing benthic foraminifera (>4% in at least one association) in each faunal association.

Fig. 7. Two-dimensional ordination of benthic foraminiferal faunal samples from the Auckland (prefixed by A) and Campbell (prefixed by C) islands, produced by Detrended Canonical Correspondence Analysis using the census data of species with >5% relative abundance in at least one sample. The common species (>5% in at least one sample, three-letter abbreviations) are plotted. Vector axes (arrows) show the correlation of faunal distribution patterns with some environmental and diversity factors. The sample associations (1–10) are those identified from the cluster analysis (Fig. 2). Species abbreviations: Asn, Astrononion novozealandicum; Bos, Bolivina subexcavata; Cac, Cassidulina carinata; Cbd, Cibicides bradyi; Egs, Eggerelloides scaber; Eiv, Eilohedra vitrea; Ela, Elphidium advenum; Gav, Gavelinopsis praegeri; Hay, Haynesina depressula; Hpm, Haplophragmoides manilaensis; Hpw, Haplophragmoides wilberti; Mis, Miliolinella subrotundata; Nf, Nonionellina flemingi; Noa, Notorotalia aucklandica; Pab, Paratrochammina bartrami; Pir, Pileolina radita; Qin, Quinqueloculina incise; Qse, Quinqueloculina seminula; Qso, Quinqueloculina suborbicularis; Qsp, Quinqueloculina subpolygona; Rob, Rosalina bradyi; Roi, Rosalina irregularis; Sie, Sigmoilopsis elliptica; Spb, Spiroplectammina biformis; Tra, Trifarina angulosa; Txe, Textularia earlandi.

1. High tidal. In common with other tropical–subpolar are recorded here only from the head of Perseverance regions, high tidal salt marsh and salt meadow environments Harbour, Campbell Island, similar high tidal environments are dominated by agglutinated foraminiferal associations were not sampled on the Auckland Islands and they will (1, 2), whereas most other shelf and bathyal environments undoubtedly be present there. have calcareous-dominated faunas (e.g. Murray, 1991; 2. Restricted bay head. This shallow subtidal association (3) is Hayward et al., 1999). Although agglutinated associations unusual and unique in the New Zealand region, where

135 B. W. Hayward et al.

subtidal substrates dominated by agglutinated foraminifera elliptica, Bolivina and a diversity of unilocular species. Do these (usually M. fusca and Trochamminita salsa) are restricted small specimens live in situ in these sheltered harbour settings, or to low salinity environments at the head of estuaries have they been winnowed out of seafloor sediment by strong (Hayward et al., 1999). Here in Tucker Cove at the head of currents or large Southern Ocean swells outside the harbour and Perseverance Harbour there is sometimes a thin low salinity transported into the harbours in suspension with the incoming surface layer, but subtidally salinity is only slightly lower tide or onshore winds? This latter method of displacement has than normal marine values. Here the water is commonly been documented in the English Channel (Murray et al., 1982) tannin-coloured from all the freshwater seepage from the and elsewhere around New Zealand (Reid & Hayward, 1997). peat soils and the sediment on the subtidal floor of the cove The strongest hint that at least some of these small light- is chocolate brown and anaerobic. Anaerobic, acidic con- weight specimens have been transported in, comes from the size ditions are inferred, which might favour this agglutinated range of C. carinata. In situ populations contain a mix of assemblage and dissolve the shells of any calcareous taxa. juveniles and adults (up to 400 µm), but in these harbour Partly dissolved shells are frequently encountered in other samples (particularly from Campbell Island), only populations samples in Perseverance Harbour. of juveniles (<150 µm) are present. The other common species 3. Exposed coasts. The two associations (9–10) which occur in (above) represented only by small specimens do not have large relatively shallow, wave- and current-swept, sandy environ- adults and thus could be in situ or displaced. It is strongly ments around the Auckland Islands have distinctively dif- suspected that most E. vitrea, S. elliptica and many of the ferent faunal compositions than the other associations unilocular specimens have also been transported in from mid- (Figs. 6, 7). Although not recorded from Campbell Island, outer shelf depths offshore, where they are relatively abundant they may be present there as no samples were available for (Hayward et al., 2007). Thus, the coarse fraction faunal com- study from similar environments around this island. positions (dominated by E. advenum, N. flemingi, N. aucklandica 4. Sheltered harbours. Five calcareous-dominated associations and Haynesina depressula) are considered to reflect more (4–8) occur within the glacially incised harbours of the correctly the in situ fauna of the sheltered harbours. two island groups. All share in common the presence of Elphidium advenum and Notorotalia aucklandica, but the Biogeographical affinities with other subantarctic island groups common co-occurrence of Nonionellina flemingi separates It is not possible to make detailed comparisons of the faunal out associations 4 and 7, which are confined to the more composition with other subantarctic island groups beyond the sheltered southern and central harbours of the Auckland SW Pacific, because the taxonomic concepts and names used in Islands at depths of 17–80 m. The faunas in muddy sand at earlier studies (e.g. Chapman, 1909; Heron-Allen & Earland, 0–40 m depth in Perseverance Harbour, Campbell Island 1932; Earland, 1933, 1935) are incompatible with this study. A (associations 5, 8) differ from those in similar environments full and consistent review of the fauna from all islands is in the Auckland Islands by the abundance of Cassidulina required for detailed biogeographical analysis and this is well carinata (faunas >63 µm) or E. advenum (>150 µm). In the beyond the scope of this study. It is possible, however, to make relatively shallow depths (10–17 m) near the more exposed some general comparisons. mouth of Port Ross, northern Auckland Islands, N. In terms of species richness, the relatively low diversity of the aucklandica reaches its greatest relative abundance and inner-mid shelf (<80 m) benthic foraminiferal fauna (including characterizes the separate association 6 (Fig. 6). uniloculars) of the Auckland Islands, 51( S (130 spp.) and The vectors on the DCCA ordination (Fig. 7) indicate a Campbell Island, 52( S (71 spp.) is of a similar order to that strong correlation of the distribution of faunal assemblages and recorded from the same depth range around South Georgia, 54( species with depth (long vector arrow), with depth increasing S (60 spp. from three samples; Earland, 1933), Tierra del Fuego, from bottom right to top left. This trend parallels the expected 53( S (75 spp. from one sample; Heron-Allen & Earland, 1932) increase in planktic foraminiferal percentage with increasing and Isla de los Estados, 54( S (76 spp. from 36 samples; distance from shore (Hayward et al., 1999) and also an unusual Thompson, 1978). The Auckland and Campbell Islands have a trend of decreasing evenness (E) in faunal structure. Both species significantly higher species richness from that recorded around richness and absolute abundance of benthic foraminifera (speci- Macquarie Island, 54( S (19 spp. from one sample; Parr, 1950), mens g1 sediment) show an increasing trend with increasing Kerguelen Islands, 50( S (33 spp. from 13 samples; Parr, 1950) water depth (Fig. 7). The short vector for sand percent- and Falkland Islands, 52( S (33 spp. from ten samples; Brady, age indicates that grain size has little influence on faunal 1884; Pearcy, 1914; Cushman & Parker, 1931; Heron-Allen & composition. Earland, 1932). The species lists for all these islands are undoubtedly incomplete and would be expanded with more ex- In situ versus transported specimens haustive sampling. The main conclusion that can be drawn, Most faunas picked from the coarser fraction (>150 µm) cluster however, is that species richness at inner-shelf depths around the separately (assocs 5, 6) from the faunas from the total sand subantarctic islands is relatively low, akin to that around the fraction (>63 µm). The coarse fraction faunas have lower species shores of Antarctica (e.g. Stockton, 1973; Delaca & Lipps, 1976; richness (mean =2.4–2.9) than the total faunas (mean =4.1– Gaz´dzicki & Majewski, 2003; Majewski, 2005), but considerably 6.9) but all have a similar range of dominance and evenness lower than around southern temperate, subtropical and tropical (Table 4). Thus, the main difference between the two size islands (e.g. Murray, 1991; Hayward et al., 1999). fractions is the additional presence in the total faunas of The larger subantarctic islands (Auckland, Campbell, numerous small, low density shells of C. carinata, E. vitrea, S. Kerguelen, South Georgia, Falklands, Estados, Fig. 8) all have

136 Benthic foraminifera of Auckland and Campbell Islands

there has never been a shallow-water connection (<500 m) between the islands or across to mainland New Zealand that would have aided progressive dispersal of shallow-water benthic foraminifera. Thus, all the intertidal and shallow-marine foraminifera that now live around the Auckland and Campbell Islands must have arrived by dispersal across the Southern Ocean. Only one species (Notorotalia aucklandica) has inhabited these island shores for sufficiently long to have evolved into a distinct endemic species. Some species that do not occur around mainland New Zealand, may have reached these islands after long journeys in the Subantarctic Current from far distant lands, such as Kerguelen Islands or even South America. The majority of species (c. 70–75%) share their geographical distribution with mainland New Zealand and show that this is by far the most usual dispersal route (possibly in both directions). The route from mainland New Zealand to the Auckland and Campbell Islands is contrary to the main net surface current direction of the Subantarctic Current (SW to NE). Surface currents commonly have large mobile eddies that at certain times could sweep larval stages, juveniles or even floating logs from southern New Zealand down to the islands, or the reverse. The best evidence that this happens comes from the shared distribution patterns themselves. The most likely source for many of the dispersed foraminifera is the southwest coast of the Fig. 8. Major island groups in the subantarctic zone of the Southern South and Stewart Islands, where currents associated with the Ocean. Subtropical Front sweep southeastwards in the direction of the Auckland Islands before swinging northeast in the vicinity of similar nearshore topography, with a mix of exposed rocky the Snares Islands (Fig. 1). This would have been particularly coasts and more sheltered, glacially carved harbours. At the emphasized during glacial periods when sea-level was c. 100 m generic level their shallow-water foraminiferal faunas have con- lower than present. siderable commonality (Heron-Allen & Earland, 1932; Earland, During the last 2–3 million years, sea surface temperatures 1933; Parr, 1950; Thompson, 1978; this study). The more around the Auckland and Campbell Islands have seldom, if sheltered harbours have a dominance of Elphidium, Cassidulina, ever, been any warmer than they are today. During the many Haynesina depressula, Nonionella-Nonionellina and the Southern glacial periods, sea surface temperatures could have been 2–5( C Hemisphere-endemic genus Notorotalia. In all places, Trifarina colder than today (e.g. Weaver et al., 1998) and could poten- angulosa becomes common in deeper parts of the inner shelf tially have killed-off island populations of some intertidal and (>30 m). In sandier, more exposed locations the faunas are shallow-marine species. A further contributor to a hypothesized usually dominated by Cibicides, Rosalina, Quinqueloculina, lower diversity foraminiferal fauna during glacials, could have Pileolina–Glabratella and Miliolinella subrotunda. Numerous, been the much lower sea-levels, when there is unlikely to have small, unilocular foraminifera have been recorded from the been many sheltered shallow-water environments similar to the shallows around all these island groups. A number of rarer present fiord-like harbours. Thus, it is suggested that a sizeable species appears to be common to all of the larger subantarctic proportion of the present shallow-water benthic foraminiferal islands, e.g. Patellina corrugata, Cornuspira involvens, Tubinella fauna of the islands has probably arrived by dispersal from funalis. Ammonia and Pyrgo appear to be absent from all except mainland New Zealand and recolonized their shores only since South Georgia (Heron-Allen & Earland, 1932) and Isla de los the Last Glacial peak, 18 000 years ago. Estados (Thompson, 1978). High-tidal marsh faunas have not been reported from most other subantarctic islands. The marsh fauna from Campbell ACKNOWLEDGEMENTS Island, 52( S, 3 spp. (associations 1, 2), is of similar low diversity The authors thank Lance Shaw and the crew of the MV to that recorded from salt marshes on Tierra del Fuego, 52( S, Breaksea Girl for field assistance during BWH’s 2004 visit to the 5 spp. (Scott et al., 1990). They have two species in common islands; New Zealand’s Institute of Geological and Nuclear (H. manilaensis, M. fusca) although the dominants differ (Tro- Science, National Institute of Water and Atmosphere and chammina inflata, Jadammina macrescens at Tierra del Fuego). Auckland War Memorial Museum for the loan of additional samples; Ewen Cameron for identifying the salt marsh grass; the Dispersal and geological history Research Centre for Surface and Materials Science, University The Campbell Plateau has been submerged continuously since at of Auckland, for the use of their SEM for photomicrography; least the Eocene, 35 million years ago, well before the eruption and Marty Buzas and Andy Gaz´dzicki for their helpful reviews. and formation of the volcanic Auckland and Campbell Islands This study was funded by the New Zealand Foundation for (Adams et al., 1979; Adams, 1983; Cook et al., 1999). Therefore, Research, Science and Technology.

137 B. W. Hayward et al.

APPENDIX A: Foraminiferal list Appendix A provides a taxonomic list of all foraminifera recorded here from the Auckland (A), Campbell (C) and Snares (S) Islands, shallower than 100 m, with references to easily accessible published figures that best portray the species as interpreted by us. Generic classification largely follows Loeblich & Tappan (1987), with the ordinal classification after Loeblich & Tappan (1992). The original descriptions of these species can be found in the Ellis & Messina world catalogue of foraminiferal species on www.micropress.org.

APPENDIX A.1: Benthic foraminifera Order Astrorhizida Lagenammina difflugiformis (Brady, 1879); Jones, 1994, pl. 30, figs 1–3. A, C Rhizammina algaeformis Brady, 1879; Jones, 1994, pl. 28, figs 1–11. C Order Lituolida Cribrostomoides jeffreysi (Williamson, 1858); Hayward et al., 1999, pl. 1, figs 23–24. A, C Glomospira cf. fijiensis Brönnimann, Whittaker & Zaninetti, 1992; Hayward et al., 1999, pl. 1, figs 3–4. C Haplophragmoides manilaensis Andersen, 1953; Edwards et al., 2004, pl. 1, figs 5–6. C Haplophragmoides wilberti Andersen, 1953; Hayward et al., 1999, pl. 1, figs 25–26. A, C Labrospira spiculolega (Parr, 1950); Parr, 1950, pl. 5, figs 8–10 (as Trochammina). A Miliammina fusca (Brady); Hayward et al., 1999, pl. 1, figs 5–6. C Reophax nodulosus Brady, 1879; Jones, 1994, pl. 31, figs 6–9. C Reophax subfusiformis Earland, 1933; Hayward et al., 1999, pl. 1, figs 15–16. C Scherochorella moniliforme (Siddall, 1886); Hayward et al., 1999, pl. 1, figs 13–14. A Spiroplectammina biformis (Parker & Jones, 1865); Jones, 1994, pl. 45, figs 25–27. C Order Trochamminida Paratrochammina bartrami (Hedley et al., 1967); Hayward et al., 1999, pl. 2, figs 1–3. A, C, S Portatrochammina sorosa (Parr, 1950); Hayward et al., 1999, pl. 2, figs 4–5. A, C Rotaliammina ochracea (Williamson, 1858); Thomas et al., 1990, pl. 1, fig. 10 (as Trochammina). A Trochammina inflata (Montagu, 1808); Hayward et al., 1999, pl. 2, figs 6–8. A Order Textulariida Eggerelloides scaber (Williamson, 1858); Jones, 1994, pl. 47, fig. 15–17. A, C Textularia earlandi Parker, 1952; Hayward et al., 1999, pl. 2, figs 22–23. C Verneuilinulla advena (Cushman, 1922); Loeblich & Tappan, 1994, pl. 19, figs 8–9. A Order Spirillinida Patellina corrugata Williamson, 1858; Hayward et al., 1999, pl. 3, figs 11–13. A, S Patellinoides conica Heron-Allen & Earland, 1932, pl. 13, figs 26–29. A Spirillina obconica Brady, 1879; Jones, 1994, pl. 85, figs 6–7. A Spirillina vivipara Ehrenberg, 1843; Hayward et al., 1999, pl. 3, fig. 7. A, S Order Miliolida Cornuspira involvens (Reuss, 1850); Hayward et al., 1999, pl. 3, fig. 16. A, C Miliolinella subrotunda (Montagu, 1808); Hayward et al., 1999, pl. 3, fig. 25. A, C Quinqueloculina bicostoides Vella, 1957; Hayward et al., 1999, pl. 4, figs 15–17. A, C Quinqueloculina delicatula Vella, 1957; Hayward et al., 1999, pl. 4, figs 23–24. A Quinqueloculina incisa Vella, 1957; Hayward et al., 1999, pl. 4, figs 25–26. A Quinqueloculina oblonga (Montagu); Yassini & Jones, 1995 (as Triloculina), figs 188–192, 196–197. A, S Quinqueloculina seminula (Linnaeus); Hayward et al., 1999, pl. 5, figs 9–10. A, C, S Quinqueloculina suborbicularis d’Orbigny, 1826; Hayward et al., 1999, pl. 5, figs 6–8. A, S Quinqueloculina subpolygona Parr, 1945; Hayward et al., 1999, pl. 5, figs 11–13. A, S Sigmoilopsis elliptica (Galloway & Wissler, 1927); Hayward et al., 1999, pl. 5, figs 16–18. A, C Tubinella funalis (Brady, 1884); Jones, 1994, pl. 13, fig. 6–11. A Order Lagenida Amphicoryne separans (Brady, 1884); Loeblich & Tappan, 1994, pl. 127, fig. 1–18. A Astacolus crepidulus (Fichtel & Moll, 1798); Hayward et al., 1999, pl. 6, figs 28–29. A Astacolus insolutus (Schwager, 1866); Hayward et al., 1999, pl. 6, fig. 30. A Astacolus vellai Saidova, 1975, pl. 50, fig. 10. A Favulina hexagona (Williamson, 1848); Hayward et al., 1999, pl. 8, fig. 2 (as Oolina hexagona). A, C Favulina punctatiformis (McCulloch, 1977); McCulloch, 1977, pl. 54, fig. 18 (as Lagena). A Fissurina baccata (Heron-Allen & Earland, 1922); Hayward et al., 1999, pl. 7, fig. 15. A

138 Benthic foraminifera of Auckland and Campbell Islands

Fissurina crucifera McCulloch, 1977, pl. 58, fig. 2. A Fissurina evoluta McCulloch; Hayward & Grace, 1981, p. 50, fig. 5e. A Fissurina evolutiquetra McCulloch, 1977, pl. 58, figs 21, 24, 25, 27. A Fissurina lucida (Williamson, 1848); Hayward et al., 1999, pl. 7, figs 20–21. A, C Fissurina marginata (Montagu, 1803); Hayward et al., 1999, pl. 7, figs 22–23. A Fissurina marginatoperforata (Seguenza, 1880); Yassini & Jones, 1995, figs 411, 413–5. A, C Fissurina staphyllearia Schwager, 1866; Sidebottom, 1912, pl. 17, fig. 20. A Galwayella trigonomarginata (Balkwill & Millett, 1884); Jones, 1994, pl. 61, figs 12–13. A Grigelis orectus Loeblich & Tappan, 1994; Hayward et al., 1999, pl. 6, figs 14–15. Guttulina irregularis (d’Orbigny, 1846); Hayward et al., 1999, pl. 7, figs 10–11. A Guttulina yabei Cushman & Ozawa, 1929; Hayward et al., 1999, pl. 7, fig. 12. A, S Homalohedra liratiformis (McCulloch, 1977); McCulloch, 1977, pl. 53, figs 25, 32. A Laevidentalina neugeboreni (Schwager, 1866); Hayward et al., 1999, pl. 6, figs 16–17 (as L. bradyensis). A Lagena crenata Parker & Jones, 1865; Jones, 1994, pl. 57, figs 15, 21. A Lagena hispida Reuss, 1858; Hayward et al., 1999, pl. 7, figs 1–2. C Lagena laevicostatiformis McCulloch, 1981; Hayward et al., 1999, pl. 7, fig. 3. A, C Lagena lyellii (Seguenza, 1862); Cushman, 1923, pl. 6, fig. 3. A Lagena spicata Cushman & McCulloch; Jones, 1994, pl. 58, figs 4, 5–6. A Lagena spiratiformis McCulloch, 1981; Hayward et al., 1999, pl. 7, figs 6–7. A Lagena sulcata (Walker & Jacob, 1798); Jones, 1994, pl. 57, figs 23, 25–27, 33, 34. A Lagenosolenia confossa McCulloch, 1977; Hayward et al., 1999, pl. 7, figs 27–28. A Lagenosolenia laciniosa Loeblich & Tappan, 1994, pl. 161, figs 11–15. A Lagenosolenia neosigmoidella McCulloch, 1977, pl. 51, fig. 9. A Lagenosolenia rara McCulloch, 1977, pl. 52, fig. 3. A Lagenosolenia strigimarginata Loeblich & Tappan, 1994, pl. 161, figs 9–10. A Lagnea sp. A Laryngosigma hyalascidia Loeblich & Tappan, 1953, pl. 15, figs 6–8. A Laryngosigma williamsoni (Terquem, 1878); Yi & Jones, 1995, figs 661–663. A Lenticulina australis Parr, 1950; Hayward et al., 1999, pl. 6, figs 31–32. A, C, S Neolingulina viejoensis McCulloch, 1977, pl. 49, fig. 8. A Nodosaria nebulosa (Ishizaki, 1943); Ishizaki, 1943, pl. 10, figs 5, 7–8 (as Lagenonodosaria). A Oolina borealis Loeblich & Tappan, 1954; Hayward et al., 1999, pl. 8, fig. 1. A, C, S Oolina globosa (Montagu); Yassini & Jones, 1995, p. 112, figs 369–370. A Oolina lineata (Williamson); Jones, 1994, pl. 57, fig. 13. A Oolina melo d’Orbigny, 1839; Hayward et al., 1999, pl. 8, fig. 3. A, C, S Parafissurina basispinata McCulloch, 1977, pl. 72, figs 1–3. A Parafissurina scaphaeformis Parr, 1950, pl. 10, figs 1–3. A Parafissurina unguis (Heron-Allen & Earland, 1913); Heron-Allen & Earland, 1913, pl. 7, figs 1–3. A Procerolagena meridonalis (Wiesner, 1931); Jones, 1994, pl. 58, fig. 19. A Procerolagena multilatera (McCulloch, 1977); McCulloch, 1977, pl. 50, fig. 5 (as Lagena). A Pseudolingulina bradii (Silvestri, 1903); Jones, 1994, pl. 65, fig. 16 (as Frondicularia). A Pseudosolenia wiesneri (Barker, 1960); Jones, 1994, pl. 59, fig. 23. A Pyramidulina n. sp. Hayward et al., 1999, pl. 6, figs 25–26 (as Pyramidulina perversa). A Sigmoidella pacifica Cushman & Ozawa, 1928; Loeblich & Tappan, 1994, pl. 149, figs 1–9. A

Order Robertinida Cerobertina tenuis (Chapman & Parr, 1937); Finlay, 1938, pl. 11, ﬁgs 4–5. A

Order Buliminida Abditodentrix pseudothalmanni (Boltovskoy & Guissani de Kahn, 1981); Loeblich & Tappan, 1994, pl. 218, figs 1–2. A, C Bolivina cacozela Vella, 1957; Hayward et al., 1999, pl. 8, figs 8–9. A, C Bolivina compacta Sidebottom, 1905; Hayward et al., 1999, pl. 8, figs 10–11. A, C Bolivina neocompacta McCulloch, 1981; Hayward et al., 1999, pl. 8, figs 12–13. A, C Bolivina pygmaea (Brady, 1881); Jones, 1994, pl. 53, figs 5–6 (as Brizalina). A, C Bolivina robusta Brady; Jones, 1994, pl. 53, figs 7–9. C Bolivina spathulata (Williamson, 1858); Hayward et al., 1999, pl. 8, fig. 17. A, C, S Bolivina subexcavata Cushman & Wickenden, 1929; Hayward et al., 1999, pl. 8, fig. 22. A, C, S Bolivina translucens Phleger & Parker, 1951; Loeblich & Tappan, 1994, pl. 213, figs 9–14. A, C Bulimina elongata d’Orbigny, 1826; Hayward et al., 1999, pl. 9, figs 6–7. A, C

139 B. W. Hayward et al.

Bulimina marginata acanthia Costa, 1856; Hayward et al., 1999, pl. 9, figs 16–17. A, C Bulimina marginata marginata d’Orbigny, 1826; Hayward et al., 1999, pl. 9, figs 13–15. A, C, S Buliminella elegantissima (d’Orbigny, 1839); Hayward et al., 1999, pl. 9, figs 18–19. A, C Cassidulina carinata Silvestri, 1896; Hayward et al., 1999, pl. 8, figs 23–24. A, C, S Cassidulina norvangi Thalmann, 1952; Nomura, 1983, figs 45–46, pl. 4, figs 12a–c, 13. C Evolvocassidulina orientalis (Cushman, 1922); Hayward et al., 1999, pl. 8, fig. 28. A Fursenkoina cf. riggii (Boltovskoy, 1954); Boltovskoy et al., 1980, pl. 34, figs 19–22. A, C Globocassidulina canalisuturata Eade, 1967; Hayward et al., 1999, pl. 8, figs 29–30. A, S Globocassidulina elegans (Sidebottom, 1910); Loeblich & Tappan, 1994, pl. 223, figs 1–6. A Globocassidulina subglobosa (Brady, 1881); Jones, 1994, pl. 54, figs 17a–c. C Rutherfordoides rotundata (Parr, 1950); Jones, 1994, pl. 52, figs 10–11 (as Fursenkoina). A Siphogenerina dimorpha (Parker & Jones); Yassini & Jones, 1995, fig. 623 (as Rectobolivina dimorpha pacifica). A, C Trifarina angulosa (Williamson, 1858) sensu lato; Hayward et al., 1999, pl. 9, figs 23–24. A, C, S Uvigerina peregrina Cushman, 1923; s.l. Jones, 1994, pl. 74, figs 24–26 (as U. bradyana). A Suborder Rotaliina Anomalinoides spherica (Finlay, 1940); Hayward et al., 1999, pl. 15, figs 27–29. A, S Anomalinoides tasmanica (Parr, 1950); Parr, 1950, pl. 15, figs 4a–c. C Astrononion novozealandicum Cushman & Edwards, 1937; Hayward et al., 1999, pl. 15, figs 8–9. A, C, S Cibicides corticatus Earland, 1934; Hayward et al., 1999, pl. 14, figs 19–21. A Cibicides dispars (d’Orbigny, 1839); Hayward et al., 1999, pl. 14, figs 22–24. A, C, S Cibicides pachyderma (Rzehak, 1886); van Morkhoven et al., 1986, pl. 22. A Cibicidoides bradyi (Trauth, 1918); van Morkhoven et al., 1986, pl. 30. A, C Cymbaloporetta bradyi (Cushman, 1915); Hayward et al., 1999, pl. 14, figs 28–29. A Discanomalina coronata (Parker & Jones, 1857); Jones, 1994, pl. 97, figs 1–2. C Discorbinella bertheloti (d’Orbigny, 1839); Hayward et al., 1999, pl. 14, figs 1–3. A, C, S Discorbinella complanata (Sidebottom, 1918); Hayward et al., 1999, pl. 14, figs 4–6. A, C, S Discorbinella deflata (Finlay, 1940); Hayward et al., 1999, pl. 14, figs 7–9. A Discorbinella subcomplanata (Parr, 1950); Hayward et al., 1999, pl. 14, figs 10–12. A Discorbinella timida Hornibrook, 1961, p. 116, pl. 14, figs 288, 293, 297. A, C Discorbinella vitrevoluta (Hornibrook, 1961); Hayward et al., 1999, pl. 14, figs 16–18. A Dyocibicides sp. A, C Eilohedra vitrea (Parker, 1953); Hayward et al., 1999, pl. 13, figs 14–16. A, C Elphidium advenum f. limbatum (Chapman, 1907); Hayward et al., 1999, pl. 17, figs 1–2. A, C Eponides repandus (Fichtel & Moll, 1798); Jones, 1994, pl. 104, fig. 19. C, S Gavelinopsis praegeri (Heron-Allen & Earland, 1913); Hayward et al., 1999, pl. 10, figs 15–17. A, C, S Haynesina depressula (Walker & Jacob, 1798); Hayward et al., 1999, pl. 15, figs 10–11. A, C Heronallenia lingulata (Burrows & Holland, 1895); Jones, 1994, pl. 91, fig. 3. A, C, S Heronallenia pulvinulinoides (Cushman, 1915); Hayward et al., 1999, pl. 13, figs 7–9. C, S Heronallenia unguiculata (Sidebottom, 1918); Sidebottom, 1918, pl. 6, figs 12–14. Neoconorbina terquemi (Rzehak, 1888); Jones, 1994, pl. 88, figs 5–8. A Nonionellina flemingi (Vella, 1957); Hayward et al., 1999, pl. 15, figs 14–15. A, C Nonionoides turgida (Williamson, 1858); Hayward et al., 1999, pl. 15, figs 16–17. A Notorotalia aucklandica Vella, 1957; Hayward et al., 1999, pl. 16, figs 13–15. A, C Notorotalia depressa Vella, 1957; Hayward et al., 1999, pl. 16, figs 16–18. C Oriodorsalis umbonatus (Reuss, 1851); Hayward et al., 1999, pl. 15, figs 24–26. C, S Pileolina radiata Vella, 1957; Hayward et al., 1999, pl. 12, figs 13–15. A, S Planodiscorbis rarescens (Brady, 1884); Jones, 1994, pl. 90, figs 2, 3. A Planoglabratella opercularis (d’Orbigny, 1826); Hayward et al., 1999, pl. 13, figs 1–3. A, C, S Rosalina bradyi (Cushman, 1915); Hayward et al., 1999, pl. 11, figs 1–3. A, C, S Rosalina irregularis (Rhumbler, 1906); Hayward et al., 1999, pl. 11, figs 4–5. A, C Rosalina paupereques Vella, 1957; Hayward et al., 1999, pl. 11, figs 6–8. A, S Rosalina vitrizea Hornibrook, 1961; Hayward et al., 1999, pl. 11, figs 9–11. A Sphaeroidina bulloides d’Orbigny, 1826; Hayward et al., 1999, pl. 11, figs 15–16. A, C, S Stomatorbina concentrica (Parker & Jones, 1864); Hayward et al., 1999, pl. 10, figs 7–8. A, S

APPENDIX B: Auckland and Campbell Islands station data Table B1 provides the data from the Auckland and Campbell Islands stations. Note that samples with a ‘c’ suﬃx represent faunal picks of the >150 µm fraction only.

140 Benthic foraminifera of Auckland and Campbell Islands

No. Station No. Cat No. Latitude Longitude Water depth Plank Alpha H E ((#S) ((#E) (m) % >63 µm >63 µm >63 µm

Auckland Islands A1 AU18098 50 30.2 166 17.0 8 16 6.7 2.5 0.54 A2 AU18099 50 31.3 166 13.6 10 19 5.8 2.0 0.35 A3 AU18100 50 31.3 166 13.7 13.5 7 3.5 1.9 0.46 A4 AU18101 50 31.3 166 13.8 16.5 27 4.7 1.5 0.25 A5 AU18102 50 50.8 166 00.9 22 2 2.3 1.4 0.40 A6 AU18103 50 50.9 166 00.9 4 13 5.2 2.1 0.42 A7 AU18104 50 51.0 166 01.0 51 11 4.8 1.8 0.35 A8 AU18105 50 48.8 166 04.6 17.5 1 2.5 1.6 0.43 A9 L12457 AU18455 50 49.8 165 55.7 9 0 2.5 1.0 0.24 A10 J531 AU18456 50 51.0 166 06.0 46 12 6.4 2.4 0.51 A11 J526 AU18457 50 50.89 166 06.0 59 23 7.1 2.3 0.40 A12 J522 F202511 50 52.5 166 09.2 80 22 6.9 2.4 0.43 A13 J515 AU18458 50 42.85 166 07.3 54 9 10.4 2.8 0.52 A14 J512 AU18459 50 41.3 166 11.5 59 28 7.2 2.4 0.47 A15 L12454a AU18460 50 29.1 166 18.2 H.T. 0 3.8 1.9 0.45 A16 L12454w AU18461 50 29.1 166 18.2 H.T. 4 2.7 1.5 0.44 A17 F201002 50 49 166 04 12 A18 F201162 50 49 166 04 18 A19 L6184 L6184 50 48.7 166 04.3 3 A20 F201106–7 50 30.5 166 17 12 Campbell Island C1 AU18107 52 33.1 166 09.3 21 45 4.8 1.4 0.23 C2 AU18108 52 33.1 166 09.8 30 42 3.1 1.4 0.32 C3 AU18109 52 32.8 166 09.5 22 5 2.9 1.3 0.30 C4 AU18110 52 32.9 166 10.3 31 55 4.9 1.6 0.28 C5 AU18111 52 33.2 166 10.1 28 38 4.1 1.5 0.29 C6 AU18112 52 33.3 166 09.5 17 50 2.2 1.1 0.31 C7 AU18113 52 33.1 166 09.1 4 56 3.7 1.4 0.30 C8 AU18114 52 33.2 166 08.8 4 45 2.3 1.5 0.49 C9 AU18115 52 33.3 166 09.0 12 59 2.8 1.2 0.29 C10 AU18116 52 33.5 166 08.6 10 64 4.5 1.5 0.26 C11 AU18120 52 33.1 166 08.7 M.T. 1 1.8 1.1 0.43 C12 B189 AU18462 52 33.3 166 09.2 19 45 3.2 1.5 0.34 C13 D29 AU18463 52 33.3 166 10.8 18 62 6.0 1.7 0.25 C14 D30 AU18464 52 33.5 166 14.0 40 47 4.8 1.5 0.24 C15 AU18121 52 33.0 166 08.7 MHWS 0 2.6 0.6 0.94 C16 AU18122 52 33.0 166 08.7 EHWS 0 0.6 0.8 0.77 C17 AU18123 52 33.0 166 08.7 MHWS 0 0.6 1.0 0.94 C18 AU18124 52 33.0 166 08.7 MHW 0 0.6 0.4 0.41 C19 AU18125 52 33.0 166 08.7 MHW 0 0.6 0.2 0.39 C20 AU18126 52 33.0 166 08.7 MHWN 0 0.4 0.1 0.56 C21 AU18127 52 33.0 166 08.7 MHWN 0 0.6 0.7 0.70

Alpha is the Fisher Alpha Index (number of species standardized by number of individuals counted). H is the Information Function (a combination of the number of species present and, to a lesser extent, the evenness of species counts). E is the Evenness (a measure of dominance versus evenness of species counts). EHWS, Extreme high water spring level; MHW, mean high water; MHWN, mean high water neap; MHWS, mean high water spring level; M.T., mid-tide level Table B1. Data from the Auckland and Campbell Islands stations

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